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L. O. HOWARD. F-ntomoloB-Ul •>. id< !.. -f..| [iurrau. 



I). B. CASTEEL, Pit. D., 
•• r: Adju Zoology, 



L. O. Howard, Entomologist and Chief of Bureau. 

C. L. Mari.att, Entomologist and Acting Chief in Absence of Chief. 

R. S. Clifton, Executive Assistant. 
W. P. Tastf.t, Chief Clerk. 

F. H. Chittenden, in charge of truck crop and stored product insect investiga- 

A. D. Hopkins, in charge of forest insect investigations. 
W. D. Hunter, in charge of southern field crop insect investigations. 
' M. Websteb, in charge of cereal and forage insect investigations. 
-v. L. Quaintance, in charge of deciduous fruit insect investigations, 
vl F. Phillips, in charge of bee culture. 

I>. M?. rtO&EBS, in charge of preventing spread of .moths, field work. 
Rolla P. Currie, in charge of editorial work. 
Mabel Colcord, in charge of library. 

Investigations in Bee Culture. 

B. P. Phillips, in charge. 

G. P. White, J. A. Nelson, experts. 

G. S. Demutii, A. H. McCbay, X. E. McIndoo, apicultural assistants. 

D. B. Casteel, collaborator. 
Pearle II. Gabrison, preparator. 

ADDITIONAL COPIES of this publication 
Xi. may be procured from the Superintend- 
ent of Documents, Government Printing 
OHice, Washington, D. C, at 5 cents per copy 

Circular No. 161. 

I nitcd States Department of Agriculture, 

L. O. HOWARD, Entomologist and Chief of Bureau. 



By D. B. Oasti i i.. I'll. l>. 

Collaborator; Adjunct Professor <>f Zoology, University of Texas, 

I n 1 1:< ©1 CTI< >N. 

Tlu" particular form of bee activity with which this paper deals 
is that which results in the removal of the wax scales from the bodies 
of the worker bees and in the application to the comb of the wax 
thus obtained. A detailed presentation of tin- facts will he given 
ami attention called to certain current conceptions of this process 
which arc in error. 

Since the bee IS a very lively Lnsecl it IS not surprising that the 
bodily movements upon which some of it- activities depend are 
extremely difficult to follow ami may easily he misunderstood. All 
of its highly specialized legs may he u-ed at once in the performance 
of some intricate process, ami the observer is in need of keenness of 
-~ i ltIi I and patience if he would gain more than an approximate 
understanding of the parts taken by tlu 1 several members. 

In the more permanent literature of apiculture and of zoology 

will he found well-written account- of the habits of lice-, accounts 

which are founded upon a large amount of careful observation and 
which represent the work of many students of bees from the time 
of lluher on. As the years have gone by, errors of sight and of 
judgmenl have gradually been eliminated, so that at the present time 
our knowledge of bee life, so far a- it goes, rests upon a fairly 
satisfactory foundation of authenticated facts. Yet many puzzling 
questions are still unanswered, and some supposed fact- may Mill he 

An examination of a number of bees from an active colony will 
show great variation in the appearance of the wax scales of ditf'er- 
.m.-.u.-, -dr. 101—12 1 


ent members of the colony. In some cases no scales can be observed, 
even upon dissection. In others the scales will be found to be ex- 
tremely thick and completely filling the wax pockets. Some bees 
will show scales in two or three pockets and none in others. Many 
of the workers will possess a complete supply of scales, either all 
of about the same thickness or varying considerably in thickn 
These and other diverse conditions present themselves for explana- 

The present account is particularly concerned with the manipula- 
tion of the wax scales. By what process or series of processes are 
the scales of wax removed from their pockets and added to the 
comb? That the wax which forms the comb is produced by the bees 
themselves, being elaborated within their bodies and given out in the 
form of thin plates or scales, is a fact well known to all students 
of bees; but many differences of opinion have been expressed con- 
cerning the exact method of wax manipulation. It is also well 
known that the workers of the hive perform many duties — build- 
ing the comb, gathering the stores of pollen and honey, caring for 
the brood and the queen, repairing, cleaning, ventilating, and guard- 
ing the hive — and it has been fairly well established that in some 
cases, at least, these duties vary with the age of the individual 
Avorker, although more accurate information on this point is much 
to be desired. Dreyling's 1 results, in particular, indicate that bees 
of certain ages are incapable of producing wax, since their glands 
s;re either undeveloped or atrophied. Do these bees use the wax 
secreted by others, taking it from them, manipulating it, and form- 
ing it into comb? By careful observation bees devoid of wax scales 
or with scales too thin for satisfactory removal ma}* be discovered 
working with the wax. Do these bees procure their wax from other 
workers, or are the} 7 merely reworking the wax of the comb? Upon 
each hind leg of a worker bee is located a peculiar pincers-like 
structure long known as the wax shears. Do bees really use this 
instrument in extracting the scales from the pockets, and if so, does 
the owner of the scale perform this operation, or is the scale re- 
moved by another worker? Or may it not be that the wax scale-- 
drop from their pockets when they reach a suitable thickness, and 
are salvaged by other workers and added to the comb? All of the 
above interpretations of these processes have been advanced by 
various observers. It is the object of this paper to present a true 
account of the manner in which the scales of wax are transported 
from their pockets to the comb and to point out some of the causes 
which lead to diversity in scale number and scale form. 

1 Drpylinp, I.. 1903. Ueber dip wacbbereitenden Organo dor Ilonigbiene. Zoologiscber 
Anzeiger, Vol. 26. 

Same. — 1905. Die wachboroitondon Organc bei don geselUg lebondon Iiionou. Zoolo- 
fiisclio Jahrbiicher, Abthellung fur Anatomic u. OntoRonie d. Tbloro, Vol. 22. 

M \NI1-I I \ I luN OF WW SCALES Ot INK llo.M 

i ii i u \\ii:i in < i m. (iiu; \ ss. 

Th" way in which the was Bcales arc formed, a- secretion products 
arising from the surfaces of the wax plates on the ventral side of 
the abdomen <>f the workers, has been well described by others and 
with apparent accuracy. The accounts <•! Dreyling embody the 
results of a very considerable amount of work, ami will, for the 
present, at least, It taken at their lull value. The work of Snod- 
»nbs ' upon the anatomy of the "wax plate- and wax glands may be 
relied upon. Only a brief statement will here It given of the struc- 
ture of these organs and of the manner in which the scales are 

\ \t well known, wax is produced by the worker bees only. The 
location of the wax-secreting sur- 
faces, or wax plates, may he readily 
determined by an examination of the 
ventral surface of a bee's abdomen. 
By stretching the abdomen Bomewhat 

it w ill he seen that each of the last 

four visible sternal or ventral plates 
i- divided into two regions: A pos- 
terior projecting edge which i- dis- 
tinctly hairy, and a smooth anterior 
half which is usually covered by the 

next preceding plate. Thi- anteri< r 
region is divided by a median ridge 
into two distinct, irregularly oval 
hich thus lie on either side 
of the midventral line. These area- 
arc the wax plate-, and upon them 
the wax scales are formed. Each one 
of the last four sternal plate- bears 
two wax plates, making eight in all. 
(S Bg.l.) 

The glands which -ci rite the \\;i\ 
lie on the floor of the abdomen im- 
mediately above and in contact with the wax plate-, and their 
secretion is deposited upon the externa] surfaces of the plates, exud- 
ing through the many minute pore- which perforate the plates. Upon 
coming in contact with the air the fluid wax hardens, forming a cov- 
ering over the entire outer surface of the plate, which gradually in- 
creases in thickness with the continued addition of wax through the 
pores. Tn this way the wax scales are produced, and since they are 

l'n;. i Ventral abdominal pintra 
nf -.\ w oi isected to -how 

the position of the wai plates. 

■ Bnodcnn, B r . 1810 The Anatomy of the Floney Bee, Bur. Bnt, Tech. s-r. 18, 
i s Dept Agt 


molded upon the surfaces of the eight wax plates they correspond to 
them in number and in form. 

In its natural position each wax scale lies between its wax plate 
and the overlapping edge of the next preceding sternal plate. The 
scale thus fits into a little crevice or wax pocket and is well protected 
from injury. If the bee extends its abdomen the rear edges of the 
scales can be seen protruding from their pockets, or if the scales be- 
come very thick they will push the covering sternal plates outward 
and will project from the pockets. 


The problem of wax secretion has been extensively studied by 
Dreyling, who shows that the wax glands differ markedly in struc- 
ture in bees of different ages. In the newly emerged bee the epider- 
mis which underlies the wax plate is composed of epithelial cells 
nearly cubical in form. As the bee grows older these cells become 
elongated and are separated by clear spaces, and when the bee has 
reached the height of its activity as a wax producer these gland cells 
are elongated and show liquid wax stored in the spaces between them. 
"When the wax-secreting period is over these cells degenerate, so that 
in sections through the glands of old field bees, or of bees that have 
lived over winter, the layer of cells beneath the wax plates appeai> 
greatly shrunken, and individual cells can be distinguished by their 
nuclei only. These histological data are given by Dreyling in sup- 
port of the conclusion that the secretion of wax in much more abun- 
dant at a certain period in the bee's life and that old bees and ver}' 
young bees are, as a rule, incapable of wax production. These con- 
clusions are in harmony with the practical experiences of bee keepers. 


In a study of the behavior during scale removal and wax building 
it is necessary to watch the bees while they are working naturally 
within the hive. To accomplish this, observatory hives are used in 
which glass is substituted for wood in part of the construction. 
Most of the work is done upon colonies in modified nucleus boxe> 
(fig. 2). The two sides are removed from each hive and are replaced 
with glass in the form of sliding doors, two to a side, and glass plates 
are fitted to the top. It all cases wooden shades cover both sides and 
top when the bees are not under observation. Although bees are 
somewhat disturbed when light is first admitted to the hive, they ap- 
pear to become accustomed to it and work normally unless the hive is 
left open for too long a period. 

"When a hive is well crowded with bees, and when the frames are 
widely spaced, the workers are apt to extend the comb above the level 
of the top bars of the frames until it comes in contact with the glass. 
This gives the observer an excellent opportunity to study the comb 

M wiiMi \ I [OS "i w w BGAJ is in' i ill HON1 I B •> 

workers at clo • range, and it also obviates the necessity of placing 
glass ends in the hive against which comb might be built. 

Even with the best of arrangements ii is difficult to follow 01 
the movements of the workers <luiiiiir the ad of scale removal. A 
nn aid to \ iaion :i Zeiss binocular microscope is used, the tubes being 
removed from the stand and held to the eye after the manner of :i 
By the use of this instrument a bee appears to acquire 

Fu.. 2. — Observatory hive, aw fitted with Midi.— glass doors, and tw.> p 

glass cover tiu> top. The sliding glass doors allow the obaerrer to gain access I 

i : iro:i of I comb without removing t ho glass from tli" entire Bide <>f Ihr 

hrre. Bcreens of wood cover the glass of tin- rides and t"i> when ' - nol under 

observatl >n. (Original. > 

the dimensions of a large-sized rat, and the action of its l<\L r - and 
mandibles may be followed with great precision. 

For the sake of later identification many of tin- bees are ma 
by painting different colors on their hacks, and some are numb 
Such distinctive marks make it possible to follow the actions of an 
individual 1hh> from day to day. 

The observations here recorded were made during the summer of 
1911 at the apiary of the Bureau of Entomology. 



Fig. 3. — Ventral view of a worker bee in the act of 
removing a wax scale. The two middle legs and 
the right hind leg are used for support, while the 
left hind leg removes the scale. (Original. I 

The determination of the exact method by which the wax scales 
are removed either comes as the result of prolonged and patient 

watching or is the product 
of good fortune. Long be- 
fore the observer is able to 
decide upon all of the de- 
tails of the process he 
becomes convinced that 
usually the scales are re- 
moved by the bee which 
secretes them and by this 
bee are masticated and 
added to the comb. The 
workers never assist each 
other in the process of re- 
moval, although, as will 
be mentioned later, free 
scales ma) T , in some ca>-e-. 
be handled by other 

As a rule, the scales are 
removed while the bee is 
standing on the comb or its support, and the wax thus obtained is 
applied to the comb near the place where it is removed. Since the 
whole process of removal takes place beneath the worker's body it 
can be observed most 
satisfactorily when 
the bee is seen from 
the side or when it is 
building comb against 
a glass plate. 

The posture of a 
bee in the act of re- 
moving a scale is 
rather characteristic 
and is at once recog- 
nized by one familiar 
with it. Immediately 
before the scale is to be 
removed the bee may 
be busily engaged upon the surface of the comb, plying with its man- 
dibles the wax of the scale last extracted or reshaping and polishing 
wax already deposited, its whole body somewhat agitated, moving 


4. — Side view of a worker in the same posture as 
that shown in figure :>. (Original.) 


backward and forward or from Bide to aid* as it adapts it- poaition t<> 
the work in hand. Suddenly ii> hody becomes very quiet The fore- 
and mandibles are raised from the comb, and the head is held 
with the face inclined tow 

ard (he comb. The 

leg of one side i- 
raised, and 



its Battened 


''"•■ ■"• Ventral riew of a worker bee showing the 
position of the was scale )us1 before 11 is grasped 
by tin' forelegs and mandibles. The scale is -till 
adhering t.> the spines <<( the i»«»i t.-n combs. The 
i- supported npon the two middle legs and 
up. .ii the bind leg which is n >t remoylng the scale 

Brs( tarsal segment 
planta is slipped along the 
ventral surfa< e of the ex- 
tended abdomen and comes 
in contact with the pro 
trading was scales of the 
corresponding aide (figs. 
:'> and I >. The weight of 
the bee is oow supported 
upon three legs; upon the 
middle leg of the side 
from which the scale is to 
be removed and upon the 
middle and hind legs of 
the other side. The first 

tarsal segment <d' the leg 

which is to remove the 

scale is now pressed firmly 
■gained the abdomen, and the edge of a protruding scale becomes 
engaged with it. steady, continuous pressure is now exerted both 
against the abdomen and toward the rear, with the result that the 

Scale is drawn out of its 

pocket but remain at- 
tached to the leg which 
removed it. The hind 
leg bearing the scale is 
now quickly Hexed tow- 
ard the thorax and 
head, thus carrying tin- 
scale forward under the 
hody of the hee and 

placing it in a position 

where it may he readily 

grasped by the forelegs 
or the mandibles (figs. 5 and 6). Sometimes the scale is appar- 
ently removed from the hind leg by the mandibles alone, hut Usually 
the forelegs aid in this process and also manipulate the scale while 

Bide vii-w nf ;i worker bee in the <amo pocrore 

;is thai Bhown in li.-iir.- r. (Original.) 



the mandibles are masticating it. After the scale has been thor- 
oughly masticated the wax is applied to the comb. 


A point of particular interest in the process of wax scale removal 
is that which deals with the manner in which the scale is grasped by 

the hind leg which removes it. As is 
well known, each hind leg of the worker 
bee bears a pincerslike structure — the 
so-called wax shears — located at the 
juncture of the tibia and the flattened 
first tarsal segment or planta (fig. 7). 
According to the statements of numer- 
ous writers, the wax scales are grasped 
between the edges of the supposed 
pincers formed by the pecten above and 
the auricle below, and are either snipped 
off or are held by the jaws of the 
pincers and thus drawn from the pock- 
ets. Cowan's 1 account may be given 
as typical of others which are current 
in the literature of apiculture and of 





Fig. 7. — Inner surface of the left 
hind leg of a worker bee, show- 
ing the position of a wax scale 
immediately after it has been re- 
moved from the was pocket. The 
scale has been pierced by seven 
of the spines of the pollen combs 
of the first tarsal segment or 
planta. The jaws of the so- 
called wax shears or pincers are 
formed by the pecten spines 
above and the surface of the 
auricle below. (Original.) 

The articulation of the tibia and planta 
being at the anterior angle, and the absence 
of the spur on the tibia (which only the 
honey bee does not possess) give the pecten a 
freedom of action it would not otherwise have 
and enable it to be used together with the 
auricle on the planta. which is quite smooth, 
as a true pair of pincers, and as an instru- 
ment for laying hold of the thin flakes of 
wax, and for bringing them forward to be 
transferred by the other legs to the jaws for 

As a matter of fact, the wax shears 
have. nothing whatever to do with the 
removal of the wax scales. They per- 
form an entirely different function, be- 
ing concerned with the gathering of 
pollen in a manner to be described in a 

future paper. 

In coming to the above conclusions the writer was first convinced 
that the so-called wax shears are not used in removing scales by 
Loting that the position of the tibio-tarsal joint at the time of scale 

1 Cowan, T. W., " The Honey Bee," 2d ed., London, 1904. 

MWIi'l I \lloN n| WA\ St All. S OF TH1 BON] .', Ull. 

removal is Buch as to make il impossible for the pincerslike crei 
ic» grasp the scale. Moreover, the open jaws of the shears poinl hit - 
terally and away from the ither than toward them, nor, 

indeed, is it possible for the shears to grasp even the projecting edj 

m\ f the ventral or lateral body plates and thus steady or guide 
tlu> leg as it seeks contact with the 

The transverse rows <>!' Bpines upon the planta, called pollen combs, 
iiinl not the was shears are instrumental in the removal of scales. 
Snodgrass (1910), in discussing the anatomy < > f the hind leg and 
,i~ functions, states that the wax is "poked out of" the "pockets 
by means of the Bpines on the feet '" " with the ordinary hairs or 
spines i i the tibiae or tarsi," and the same general conclusions were 
reached independently by the writer, but with this exception; only 
the spines of the first tarsal segment (planta) function in this 
manner, and usually < nly certain large spines in the rows at the dis- 
tal end of this segment 

It is exceedingly difficult to capture a bee at the very moment at 
which the scale is being drawn from its p< cket and before it has been 
carried to the month, and even if this is acc< mplished the captive is 
very likely to drop the scale from the hind leg in it- struggles to 
escape. If. however, one i> successful, the scale-removing leg will 
show the little wax scale adhering to the distal end of the inner 
surface of the first tarsal segment, being pierced in several pla 
by the stn Dg Spines which project from the lower rows of the pollen 
combs. (See liir. T.) 

It can also he shown experimentally that this method of remov- 
ing the wax scales is entirely possible, for if the hind tarsus of a bee 
i> mounted upon a small -tick an. I i- gently rubbed ah ugthe ventral 
side of a fully extended dead bee's abdomen, holding it in such a 
p' sition that the pollen combs brush over the projecting edges of the 

les, one of the scales will probably be removed and will be seen 
adhering to the spines in the manner above described. 

In any hive where comb i> being constructed rapidly many free 
scales will be found upon the bottom board and upon the lower bars 
of the frames. If these scab- are examined microscopically some 
will lie found without mark- upon them, having evidently been 
lot sened from their pockets accidentally during the movements of 
the worker- over the comb and around the hive. Others will --how 
certain marks ami scratches upon them, indicating that they were 
voluntarily removed from the picket-, and in some ca-es they may 
bear the mark- of the mandibles, showing thai they were dropped 
during the process of mastication. Most of tin scales which are 
marked at all are indented with several small puncture- showing the 
places where the spines of the pollen comb.- have pierced them. 
These .-cars are exactly similar in appearance to those on the scale 


shown in figure 7. Such free scales are not marked as they would be 
had they been extracted By such a structure as the so-called wax 

So far as can be determined there does not appear to be any regu- 
lar order for the removal of scales. One may be taken from the 
left side and then one from the light, or the bee may remove two or 
three from one side in succession. An attempt to remove a scale is 
by no means always successful, the worker often trying first one side 
and then the other, pressing the pollen combs against the more ante- 
rior scales and running them down to the most posterior, until at 
last a scale is impaled upon the spines or the bee discontinues its 


When a scale has become attached to the spines it is transferred to 
the mouth with great rapidity, so swiftly, in fact, that the e3 r e can 
scarcely follow the action. This is not surprising, for it is necessary 
only to flex the leg toward the head to bring the scale in close con- 
tact with the forelegs and mandibles. The leg is rotated through 
the arc of a circle, downward, forward, and upward, while at the 
same time the head is slightly turned under to receive the scale. 
The process of mastication is more prolonged. It is usually sup- 
posed that the pure wax of the scale differs in chemical composition 
from the wax of the comb, this change being accomplished during 
mastication, by which process the wax is mixed with saliva, becomes 
translucent rather than transparent, changes somewhat in color, and 
becomes more pliable. 

The behavior of a bee upon receiving a wax scale at its mouth is 
subject to considerable variation. On some occasions the scales are 
apparently manipulated by the mandibles alone, while at other times 
the forelegs are brought into requisition and assist the mandibles. 
"When a scale is thin and small and has been firmly grasped by the 
mandibles little assistance is needed from the legs. But if a 
scale of medium or extra thickness is presented, or if the mandibles 
do not hold it securely and it is in danger of falling from the mouth, 
the two forelegs are used to great advantage in readjusting the scale 
and in so holding it that the mandibles may be applied to it most 
advantageous!}'. If a scale is small and thin, it may be masticated 
entirely before any wax is applied to the comb; but if of considerable 
size a portion only may be prepared, this deposited upon the comb, 
and then the remainder treated in a similar manner. 

As a rule the wax which is deposited upon the comb by the pro- 
ducing bee is first subjected to the action of the mandibles and 
mixed with saliva. Such, however, is not always the case, for some 
bees appear to be " careless " and will mingle small unchewed por- 
tions of scales with the masticated wax. Indeed, it is not uncom- 

M win i \ I h>\ 0] \\\\ BCALBB OF TH1 B0VX1 BBB. 11 

mon to find Dearly perfect Bcales mixed with the wax of a newly 
made comb. The masticated wax itself is Bpongy and flaky when 
it i- deposited by the producing bee and will Later be reworl 
thereby gaining greatly in compactness and smoothne 

The entire process <>t' the removal of > ne scale, its mastication, and 
tin- application of the was to the comb is completed in :i I >« mi four 
minutes, only a very small portion of this interval being consumed in 
the work of extracting the scale frOm it- pocket and passing ii to i lu j 
mouth, except in cases in which scales appear t<> be removed with 

1 I.I I -< \|,KS. 

When wax scales are voluntarily removed they are taken ofT by 
the bee which secretes them and in the manner above described. 
Many, however, an' accidentally detached, being loosened from their 
pocket- by movements of the abdomen, incidental cleansing move- 
ments of the legs, or by contact with objects both within and without 
the hive. Such scales, and also those which are dropped in the 
course of transference from the wax pocket to the mouth, may or 

may not be recovered later and added to the comh. Since old wax 

is used over and over again in the rebuilding of comb, it i- hut 
natural to expect that scattered scales would likewise be utilized by 
the colony and not be allowed to go to Waste, and it is probably true 
that such is DSUally the ca-e. Yet there appeal's to he no concerted 

action among the workers to salvage Buch particle- of wax, no class 
of comb workers whose duty it is to pick such material from the 
hottom board of the hive and carry it to the comb. Scales which 
drop are likely to remain for a long time, and some may even he 
carried out through the entrance with waste material. If. however. 
Male- accidentally dislodged or voluntarily removed fall on the 
comb among the comb workers they are often noticed by them, picked 
up. masticated, and built into the comb. If a scale -lip- from the 
pollen comix or is fumbled by the bee before being grasped by the 
mandibles, it i- seldom recovered by the worker to which it Ixdongs 
Hide-- it fall- very near her or she -tumhle- upon it accidentally. 


Although a bee endeavor- to remove an entire wax scale at one 
operation, the attempt i- not always successful. A scale that lias 
become very thick is difficult of removal, particularly so if the outer 
edge is broken or beveled. When the hec applies its pollen combs to 

such a scale the spine- may fail to get a hold upon the wax. or they 
may not become sufficiently well fixed in it to make possible the re- 
moval of the entire male. Instead of this, shreds and small pieces of 
wax arc turn oil and remain -ticking to the bristles of the pollen 


combs. These may be entirely disregarded by the bee, or they may 
be cleaned off by scraping the combs together, the shreds of wax 
dropping to the bottom of the hive. More usually, however, if a 
worker is actively engaged in the task of adding to the comb these 
hits of wax will be carried forward to the mouth, masticated, and 

In one case which came under observation a worker had removed 
all of its wax scales except a very large, thick one which was evi- 
dently sticking tightly in its pocket. Repeated efforts were made by 
the bee to accomplish the extraction of this scale, but with only 
partial success, since the main portion of the scale remained in the 
pocket. But, as the result of its efforts the bee succeeded in beveling 
off the entire projecting edge of the scale, rasping it off bit by bit 
and carrying the small pieces forward to the mouth, masticating 
them, and dispositing the wax upon the comb. 


The presence of well-developed scales protruding from the pockets 
of a worker does not necessarily indicate that this individual will 
shortly add this wax to the comb, even though the colony may at the 
time be producing comb at a rapid rate. Such a bee may be working 
upon the comb as a molder of wax rather than as a producer. One 
who is intent upon a study of the process of scale removal will often 
be disappointed after following for a time the movements of a 
worker that is evidently manipulating wax and which shows the 
protruding edges of scales beneath its abdomen, for the wax with 
which it is working is being picked up, little bj r little, from the comb 
and comes not from its own body. This reworking of wax is one of 
the most characteristic features of comb construction, for it goes on 
continually while new comb is being produced,. and it, is, of course, a 
necessary process in the reconstruction of old comb. 

The claim has been made by several investigators and writers that 
the bees which sculpture the wax are not at the same time concerned 
with its secretion and deposition — that there are producing bees 
and building bees. In a sense this is true, but not entirely so. With- 
out doubt many active comb workers are. at the time, nonproductive, 
for the wax glands of many are not functionally active. The re- 
sults of Dreyling would indicate that the old bees, at least, might be 
considered as falling in this class, and the direct observations of the 
writer lead to the conclusion that, old bees devoid of wax scales per- 
form a considerable share of the labor of reworking newly deposited 
wax and of shaping and polishing the cells of the comb. 

However, as noted above, bees with well-developed wax scales 
often busy themselves with wax working rather than with produc- 
tion. Moreover, a bee that is removing its scales may discontinue 


this work and give its attention tu the molding of wax laid down 
l)\ others. This may occur immediately after a worker has removed 
lies, or the bee may turn to sculpturing while Beveral 
yet remain in the pockets. Ii is thus evidenl thai the produc- 
ing bee may also be a worker of wax produced by others ami that 
nonproductive bees do m>t monopolize the work of Bculpturing and 
polishing the comb. 

-I M M KB1 . 

A- is well known, the wax produced by the worker bee occurs in 
tlii> form of scales, eight in number, which appear upon the surfaces 
of (lie eight wax plates. These wax plates arc located upon the 
last i'mir visible ventral plates of a worker bee's abdomen. The wax 
reted by glands which lie upon the inner surface of each wax 
plate. The liquid wax exude- through poi ich perforate the 

Wax plate-, and it harden- to form the scales a- il conic- in contact 
with the air. 

Unless accidentally dislodged the wax scales are always removed 
and manipulated by the bee which secretes them. 

in the process of removal the scale is not grasped by the so-called 

wax shears, hut it i> pierced by a few of the -till' -pine on the distal 

end of the first tarsal segment of the hind leg and i- then drawn 

from it- pocket and remain- adhering to these -pine- until removed 

■ ia.-ticat ion. 

By flexing the hind leg the scale i- brought f< rward beneath the 
bee's body and into proximity with the mouth. In the proce 
mastication the forelegs usually aid the mandibles by holding the 
scale in an advantageous position. 

Xo definite sequence i- observed by the bee in the order in which 
it removes it- scales. 

a rule entire scales are removed at one operation, although it 

times ha] | - thai ;i thin scale is broken in extracting it from 

its pocket or an extremely thick one is gradually beveled off by the 
continued rasping of the pollen combs. 

Scales which are removed accidentally or which are dropped 
during manipulation may he recovered later and built into the i 
hut the recovery of free scales is usually not accomplished by the 
bee which secreted them. 

ich are producing wax may also rework the masticated 
W0K laid down by other-. Producing bees may turn to the work of 
building and sculpturing the comb either before all their scales 
are removed or immediately after this has been accomplished. 



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